Acta zoologica cracoviensia, 52B(1-2): 61-72, Kraków, 30 June, 2009 doi:10.3409/azc.52b_1-2.61-72

Szeptyckiana, a new of the feather mite family Pterolichidae (Acari,Astigmata,Pterolichoidea)fromtheBlack-thighed Falconet Microhieraxfringillarius (,)

Jacek DABERT and Anna LABRZYCKA

Received: 3 April 2009 Accepted: 21 April 2009

DABERT J., LABRZYCKA A. 2009. Szeptyckiana, a new genus of the feather mite family Pterolichidae (Acari, Astigmata, Pterolichoidea) from the Black-thighed Falconet Micro- hierax fringillarius (Falconiformes, Falconidae). Acta zoologica cracoviensia, 52B(1-2): 61-72. Abstract. Szeptyckiana, a new genus of the feather mite family Pterolichidae (Pterolichoi- dea), is described based on two new , S. macrosetosa sp. n. (type species), and S. microsetosa sp. n., collected from the wing feathers of museum specimens of the Black- thighed Falconet Microhierax fringillarius (DRAPIEZ, 1824) (Falconidae). This highly plesiomorphic new genus is characterized by pseudorutellar membranes with net-like pat- tern of hexagonal cells, complete idiosomal chaetotaxy, lanceolate setae c3, free epimer- ites I, and dagger-like setae cG on genua I and II. Males have very short opisthosomal lobes with tongue-shaped terminal membranes, genital and anal region not sclerotized, medium-sized aedeagus and minute adanal discs, not modified legs similar in size, tarsi IV with small apico-ventral claw, and setae d on tarsi IV shaped as short sticks; seta e ab- sent. Females have large horseshoe-shaped epigynum, 1.4-1.7 times longer than wide, and the ends of legs IV not reaching posterior rounded margin of the body. The occur- rence of two Szeptyckiana species on a single individual of M. fringillarius is the first rec- ord of feather mite lineage duplication on falconiform . Key words: Feather mites, Astigmata, Pterolichidae, falconets, new taxa.

Jacek DABERT, Department of Morphology, Faculty of Biology, Adam Mick- iewicz University, Umultowska 89, 61-614 Poznañ, Poland. E-mail: [email protected] Anna LABRZYCKA, Molecular Biology Techniques Laboratory, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznañ, Poland. E-mail: [email protected]

I.INTRODUCTION

The family Pterolichidae TROUESSART et MÉGNIN, 1884 with about 120 named genera and nearly 400 species is the largest and most diverse taxon among the feather mites. Representatives of this family are found on diverse non-passerine birds, mostly on terrestrial ones. Among hosts of these mites are also birds-of-prey (Falconiformes). However, the main part of falconiform acaro- fauna are numerous members of the family Gabuciniidae GAUD et ATYEO, 1975. The family Ptero- J. DABERT, A. LABRZYCKA 62 lichidae is poorly represented by one subfamily Pterolichinae and three genera: Epopolichus GAUD, 1981, Falcolichus GAUD et ATYEO, 1996, and Pseudalloptinus DUBININ, 1956. Two later genera arerestrictedtoFalconiformes,whereas Epopolichus issharedwithhoopoes(Upupidae). Feather mites remain in plumage also after bird’s death as dried “mummies” that gives an opportunity to examine feather mites acarofauna on preserved birds in ornithological collections (DABERT &ALBERTI 2008). During systematic studies on feather mites carried out in the ornitho- logical collections of German museums one of us (JD) found some representatives of pterolichid mites on dry skins of the two individuals of falconets, the world’s smallest falconiform birds. Sam- pled mites belong to a new genus (with two species), distinctly different to all genera known previ- ously from falconiform birds. The only feather mite species reported so far from falconets belong to two genera of the family Gabuciniidae: Hieracolichus GAUD et ATYEO, 1975 collected from Micro- hierax fringillarius (DRAPIEZ, 1824) by MÉGNIN and TROUESSART (1884) and Coraciacarus DUB- ININ, 1956 reported from M. caerulescens (L., 1758) by MCCLURE and RATANAWORABHAN (1973). However, these reports are of very limited credibility, because Coraciacarus spp. and Hieracoli- chus nisi (CANESTRINI, 1878) are taxa not found on falcons. They represent most probably either ac- cidentalcontaminationorwrongspeciesidentification.

II.MATERIALSANDMETHODS

The material used in the present study was collected in the ornithological collection of the Zoo- logical Institute and Museum, Ernst-Moritz-Arndt University, Greifswald, Germany from dry bird skins and was preserved in 70% ethanol. Before mounting, mites were softened and cleared in 10% lactic acid at +60ºC for 2-3 days. For light microscope study the mites were mounted on slides in Faure medium (EVANS 1992) and investigated under the light microscope Olympus BX51 with No- marsky differential interference contrast (DIC). Drawings were made using a camera lucida draw- ingdevice. All measurements are given in micrometers (Fm). Length of gnathosoma was measured from the distal tips of palps to the basal margin of subcapitulum. Width of gnathosoma was measured at the widest section of subcapitulum. Idiosoma length was measured from the anterior margin of pro- dorsum to the posterior end of body (except terminal membranes in males). Width of idiosoma was measured at the level of setae c2. Distances between setae were measured as the shortest distance between centers of their alveoli; for setae of different rows the measure is a mean taken from both sides of the mite. Measurements separated by dash are the min-max range. Abbreviations used in measurements and taxa descriptions: L/W – length to width ratio, P/H – propodosoma to hystero- somalengthratio. Chaetotaxy nomenclature follows that of GAUD &ATYEO (1996). Terminology of pterolichid morphology is after DABERT (2003). Scientific English and Latin names of birds as well the higher- level classification of birds are those of DICKINSON (2003). The type materials are deposited in the DepartmentofAnimalMorphology(AMU), Poznañ, Poland.

III.DESCRIPTIONSOFNEWTAXA

Szeptyckiana gen.n. T y p e s p e c i e s. Szeptyckiana macrosetosa sp. n. from Microhierax fringillarius (DRAPIEZ,1824) (Falconidae). D i a g n o s i s. Both sexes: Small-sized mites, males about 230, females about 310-330 in length. Subcapitulum of gnathosoma transversely striated, pseudorutellar membranes with hexago- nal reticulation (Fig. 6). Dorsal shields entire, well developed covering almost entire pronotum and hysteronotum. Supranal concavity well developed, circular. Idiosomal chaetotaxy complete, setae Szeptyckiana gen. n. from falconets 63 vi paired, remnants of alveoli of setae ve present. Scapular setae simple; si minute, set close to mac- rosetae se, medially. Setae c3 lanceolate. Epimerites I free; coxal fields I-II not sclerotized, III-IV with well developed shields along epimerites. Setae cG on genua I, II thickened, dagger-like. Solen- idion ó ongenuIIIpresent, ó2 ongenuIabsent. Male: Body more stocky than in females, twice longer than wide. Opisthosomal lobes very short, separated by small interlobar cleft. Lobes with tongue-shaped terminal membranes. Genital organ medium-sized; genital and anal region without sclerotized shields. Adanal discs minute, highly regressive. All legs similar in size, relatively short and not modified. Tarsi IV with small apico-ventral claw; no other apophyses present. All tarsi with well developed ambulacra. Setae d on tarsiIVshapedasshortsticks, e absent. Female: Idiosoma more slender than in males, about tree times longer than wide. Body terminus rounded with poorly expressed incision between setae h3. Dorsal opisthosoma with poorly devel- oped suture lying antero-lateraly to supranal concavity. All dorsal, lateral, and terminal setae of hys- terosoma (except h2 and h3) piliform, short. Epigynum horseshoe-shaped, not touching anterior epimerites, longer than latigenital sclerites of oviporus, 1.4-1.7 times longer than wide. Primary spermaduct of spermatheca relatively short and wide, with distal part spring-like, stretchy; secon- dary spermaducts shaped as very short, strongly sclerotized cylinders; copulatory opening set dor- sallyinsupranalconcavity(Fig.5D).LegsIV notreaching posteriormarginofthebody. D i f f e r e n t i a l d i a g n o s i s. The new genus is a highly plesiomorphic taxon devoid of clear apomorphies. That makes difficult to enumerate its specific features enabling an easy single- character distinguishing from other mites of the subfamily Pterolichinae. It is possible that the net- like pattern of pseudorutellar processes is a unique character of this genus, but temporarily this ob- servation is useless because of lacking data about shape of this structure in nearly all remaining Pterolichidae. Also the very long epigynum of female is rather unusual in pterolichins. Most genera have arched or horseshoe shaped epigynum, at most as along as wide. Females of the new genus have epigynum 1.4-1.7 times longer than wide, a situation observed also in Synapsilobus GAUD et ATYEO, 1996. But this genus in comparison to Szeptyckiana gen. n. is very different in morphology andhostrange(Galliformes). Among three genera of pterolichine mites occurring on falconiform birds two are reported from representatives of the family Falconidae: Epopolichus GAUD, 1981 (known also from Upupifor- mes) and Falcolichus GAUD et ATYEO, 1996. The third genus, Pseudalloptinus DUBININ, 1956 oc- curs on Accipitridae. However, the new genus is very different from all these genera as presented in the Table 1. The combination of these relatively not numerous characters enables also differentiat- ingthenewgenusfromallothernamedgeneraofPterolichinae. E t y m o l o g y. The new genus Szeptyckiana is dedicated to the memory of late Prof. Andrzej SZEPTYCKI,anoutstandingPolishzoologist,anhonestman,ourfriendandteacher. H o s t s. Both known species inhabits the Black-thighed Falconet Microhierax fringillarius Falconiformes,Falconidae). R e m a r k s. Occurrence of two or more closely related feather mite species on a single bird species is an effect of a duplication of the ancestral parasite line into several new species invading different discrete microhabitats on/in the plumage of a single host species (DABERT 2005). The most spectacular examples of such multiplification are parrots (Psittaciformes), where as much as 7 different Fainalges species (Xolalgidae) may occur on Aratinga cholochlora (PÉREZ 1995). The present finding of two different Szeptyckiana species inhabiting feathers of Micohierax fringillarius isthefirstreportoffeathermiteduplicationinfalconiformbirds. Szeptyckianamacrosetosa sp.n. (Figs 1, 2, 5, 6B, D) D i a g n o s i s. Setae vi in females (and probably in males) lanceolate, at least four times longer than distance between them. Terminal membranes of males tongue-shaped, as long as wide; aedea- J. DABERT, A. LABRZYCKA 64 Table1

Characters differentiating the Szeptyckiana gen. n. and known pterolichine genera offalconiformbirds(Falconiformes)

Character Szeptyckiana gen. n. Epopolichus Falcolichus Pseudalloptinus

Genital setae g present absent present present dagger-shaped or Setae c3 dagger-shaped setiform setiform setiform Solenidion ó on genu III present absent present present Epimerites I free fused free fused Setae d1 present absent present absent Setae e1 present absent present absent Setae h1 present absent absent absent Terminal membranes present absent absent present or absent (male) Paragenital apodemes (male) absent absent present absent Apical claw on tarsi IV (male) present absent present present Arched apodeme between absent absent absent present trochanters IV (male) divided into three parts or with separate with separate Hysteronotal shield (female) entire with large desclerotized pygidial part pygidial part medio-terminal area Epigynum (female) much longer than wide much shorter than wide shorter than wide much shorter than wide not reaching body extending body extending body extending body Legs IV (female) terminus terminus terminus terminus gus whip-like, distinctly wider at its base than in distal part. Distance between setae h1: h1 in fe- malesaboutthreetimeslongerthandiameterofsupranalconcavity. T y p e m a t e r i a l. From Microhierax fringillarius (type host): 1 male holotype, 3 females paratypes, Java, 1949, collector unknown, AMU G19 (Greifswald 3254). 1 female paratype, same data,AMUG20(Greifswald3256). D e s c r i p t i o n. Male, holotype (Figs 1, 5A, C, E, 6B, D): Gnathosoma with rectangular, transversely striated subcapitulum, length 40, width 38, L/W 1.1. Pseudorutellae with net-like pat- tern. Idiosoma moderately elongated, nearly rectangular, slightly narrowing terminally, length 227, width 107, L/W 2.1. Propodosoma and hysterosoma length 73 and 154, respectively, P/H 0.5. Opis- thosoma rectangular with opisthosomal lobes very short, rounded, separated by wide triangular and shallow cleft; width at level of setae ps1 14, length 4. Terminal membranes tongue-shaped, length 17, width 17. Pronotal shield entire, covering almost the whole pronotum; shield uniformly dotted with rounded latero-terminal corners and straight posterior margin. Lateral incisions around setae se poorly sclerotized. Scapular shields shaped as small sclerites dorsally, extended ventrally as oval sclerites. Humeral shields not visible dorsally. Hysteronotal shield covers the whole hysteronotum. Entire shield uniformly dotted with two irregular lacunes posterior to setae e1. Supranal concavity circularwithwellsclerotizedmargins,lyingbetweensetae h1. Szeptyckiana gen. n. from falconets 65

Fig. 1. Male of Szeptyckiana macrosetosa sp. n., ventral (left) and dorsal (right) view.

Setae vi probably thickened (only big alveoli remained on the slide). Setae si set close to se, medially. Cupulae ia with well developed surrounding sclerotization. Setae c1 set at anterior margin of hysteronotal shield. Gland opening gl set much closer to the level e1-e1 than to level d2-d2. Setae h1 set at lateral margins of supranal concavity approximately in its middlength. Setae f2, ps2 and ps1 lanceolate, f2 and ps2 subequal in length, ps1 about three times longer. Setae h2 and h3 shaped as macrosetae, subequal in length. Distance between dorsal setae: se:se 52, si:si 40, c1:c1 51, d1:d1 36, c1:d1 48, c2:c2 99, d2:d2 66, c2:d2 77, e1:e1 29, e2:e2 72, d1:e1 51, d2:e2 51, h1:h1 20, e1:h1 28, h2:h2 54, h3:h3 39, ps1:ps1 29, h1:ps1 16. Coxal fields I and II without sclerotized shields, coxal fields III and IV with well developed shields along epimerites; lateral shields of epimerites IV shaped as longitudinal rectangles, medial shields as minute triangles. Base of aedeagus located on the level of insertions of trochanters IV. J. DABERT, A. LABRZYCKA 66

Fig. 2. Female of Szeptyckiana macrosetosa sp. n., ventral (left) and dorsal (right) view.

Aedeagus 46 in length directed anteriorly and bent backwards reaching its base; aedeagus gradually narrowing, several times thinner in distal part than at the base (Figs. 5A, 6D). Genital papillae set at the level of the aedeagus bend. Setae g inserted anterior to the genital organ and distinctly posterior to the level of setae 3a, 3a posterior to 3b. Adanal discs minute, 6 in diameter, surrounded by mem- braneswithsclerotizedcentralpart. All legs similar in length and width. Tarsi IV with an apico-ventral triangular apophysis with acute apex (Fig. 5C). Legs IV reaching posterior margin of terminal membranes by ambulacra. Se- tae cG on genua I and II dagger-like, about twice shorter than the width of corresponding podomere (Fig.5E).Setae d minute,stick-like, e absent ontarsiIV. Szeptyckiana gen. n. from falconets 67

Fig. 3. Male of Szeptyckiana microsetosa sp. n., ventral (left) and dorsal (right) view.

Female, paratype (Figs. 2, 5B, D, E): Gnathosoma shaped as in males, length 48-52, width 44-49, L/W 1.1. Idiosoma length 311-324, width 118-130, L/W 2.4-2.6. Propodosoma and hystero- soma length 79-83 and 230-241, respectively, P/H 0.3-0.4. Body more elongated than in males with rounded posterior margin having poorly expressed, wide and shallow terminal cleft. Sclerotization of pronotum similar to that of males. Hysteronotal shield covers the entire dorsal hysterosoma. Poorly expressed arched suture delimits pygidial part of the shield. Hysteronotum uniformly dotted. Supranal concavity shaped and situated as in males; the copulatory opening situated close to poste- riormarginoftheconcavity(Fig.5D). Setae vi lanceolate, about four times longer than distance between them. Cupulae ia shaped as in males. Lateral setae c2, d2, e2, f2 short, piliform. Distances e1:e1 shorter than h1:h1, e1:h1 equal to h1:h1, h1:h1 three times longer than diameter of supranal concavity. Setae h2, h3 shaped as in males; ps1 hair-like, slightly longer than distance between them. Distance between dorsal setae: se:se 58-60, si:si 45-46, c1:c1 53-64, d1:d1 43-47, c1:d1 58-60, c2:c2 111-119, d2:d2 77-85, c2:d2 89-92, e1:e1 26-31, e2:e2 96-105, d1:e1 106-107, d2:e2 87-88, h1:h1 32-37, e1:h1 29-36, h2:h2 48-53, h3:h3 27-31, ps1:ps1 10-12, h1:ps1 24-27. Sclerotization of coxal fields similar to that of males. Ventral bends of hysteronotal shield form weakly sclerotized opisthoventral sclerites. Epigynum shaped as a large horseshoe-like sclerite, J. DABERT, A. LABRZYCKA 68

Fig. 4. Female of Szeptyckiana microsetosa sp. n., ventral (left) and dorsal (right) view. width 38-46, length 65-70, L/W 1.5-1.7. Tips of the epigynum branches reach the level of setae g and extend beyond the ends of latigenital sclerites (Fig. 2). Genital acetabula set posterior to setae 3a and at the level of 3b, close to epigynum branches or on these sclerites. Distance 3b:g about twice longerthan g:4a, 3a:3b approximatelyequalto 3b:g. Legs IV reaching by ambulacra the level of setae ps3. Genual setae cG I, II dagger-like, about twiceshorterthanthewidthofcorrespondingpodomere(Fig.5E). Juvenilestagesunknown. E t y m o l o g y. The specific name (macro, L. – large + saeta, L. – hair) derives from the large lanceolatesetae vi. Szeptyckiana gen. n. from falconets 69

Fig. 5. Details of morphology of Szeptyckiana gen. n. A – genital region of male, B – cupula ia of female, C – tarsus IV of male, D – spermatheca of female, stretched (left) and contracted (right), E – genu I (above) and II (below) of male (left) and female (right). ma – S. macrosetosa sp. n., mi – S. microsetosa sp. n., gp – genital papillae, hs – head of spermatheca, pa – primary spermaduct, sd – secondary sparmaduct, co – copulatory opening, sc – supranal concavity.

Szeptyckianamicrosetosa sp.n. (Figs 3, 4, 5, 6A, C)

D i a g n o s i s. Setae vi in females (and probably in males) hair-like, at most twice longer than distance between them. Terminal membranes of males tongue-shaped, twice shorter than wide; ae- deagus massive, only slightly wider at its base than in distal part. Distance between setae h1:h1 in femalesasthediameterofsupranalconcavity. J. DABERT, A. LABRZYCKA 70 T y p e m a t e r i a l. From Microhierax fringillarius (type host): 1 male holotype, 1 female paratype,Java,1949,collectorunknown,AMUG20(Greifswald3256). D e s c r i p t i o n. Male, holotype (Figs 3, 5A, C, E, 6A, C): Gnathosoma with rectangular, transversely striated subcapitulum, length 44, width 44, L/W 1.0. Pseudorutellae with net-like pat- tern. Idiosoma stocky, narrowing terminally, length 226, width 148, L/W 1.5. Propodosoma and hysterosoma length 67 and 159, respectively, P/H 0.4. Opisthosoma ovate with opisthosomal lobes very short, rounded and separated by triangular cleft; width at level of setae ps1 11, length 5. Termi- nal membranes tongue-shaped, short, length 12, width 23. Pronotal shield entire, covering almost the whole pronotum; shield uniformly dotted with rounded latero-terminal corners and straight pos- terior margin. Lateral incisions around setae se poorly sclerotized. Scapular shields shaped as small sclerites dorsally, extended ventrally as oval sclerites. Humeral shields developed dorsally as longi- tudinal sclerites. Hysteronotal shield covers the whole hysteronotum. Entire shield uniformly dotted without lacunes. Supranal concavity circular with well sclerotized margins, lying between setae h1. Setae vi in the male examined lost, but probably hair-like and short, since remained alveoli are small. Setae si set close to se, medially. Cupulae ia with well developed surrounding sclerotization. Setae c1 set on anterior margin of hysteronotal shield. Gland opening gl set closer to the level d2-d2 than to level e1-e1. Setae h1 set on lateral margins of supranal concavity close to its anterior margin. Setae f2, h2, h3, and ps2 absent on the slide; alveoli large as in S. macrosetosa. Distance between dorsal setae: se:se 70, si:si 56, c1:c1 60, d1:d1 49, c1:d1 42, c2:c2 124, d2:d2 73, c2:d2 74, e1:e1 42, e2:e2 64, d1:e1 52, d2:e2 54, h1:h1 15, e1:h1 33, h2:h2 63, h3:h3 47, ps1:ps1 36, h1:ps1 24. Coxal fields I and II without sclerotized shields, coxal fields III and IV with well developed shields along epimerites; lateral shields of epimerites IV shaped as a square, medial shields as long and narrow triangles. Base of aedeagus located between levels of insertions of trochanters III and IV. Aedeagus 64 in length, massive, only slightly thinner in distal part than at the base, directed an- teriorly and bent backwards reaching its base (Figs 5A, 6C). Genital papillae set laterally to the genital organ at the middlength of the ascended part of aedeagus. Setae g inserted laterally to the an- terior bent of the genital organ and anterior to the level of setae 3a, 3a posterior to 3b. Adanal discs minute,6indiameter,surroundedbymembraneswithsclerotizedcentralpart. All legs similar in length, legs IV slightly wider than remaining ones. Tarsi IV with an apico- ventral triangular apophysis with blunt apex (Fig. 5C). Legs IV reaching posterior margin of termi- nal membranes by distal ends of tarsi. Setae cG on genua I and II dagger-like, about 1.5 times shorter than the width of corresponding podomere (Fig. 5E). Setae d minute, stick-like, e absent on tarsiIV. Female, paratype (Figs 4, 5B, D): Gnathosoma shaped as in males, length 53, width 52, L/W 1.0. Idiosoma length 330, width 140, L/W 2.4. Propodosoma and hysterosoma length 89 and 241, re- spectively, P/H 0.4. Body more elongated than in males with rounded posterior margin having poorly expressed small terminal incision. Sclerotization of pronotum similar to that of males. Hys- teronotal covers the entire dorsal hysterosoma. Poorly expressed arched suture delimitate pygidial part of the shield. Hysteronotum uniformly dotted. Supranal concavity shaped and situated as in males;thecopulatoryopeningsituatednearthecenteroftheconcavity. Setae vi hair-like, short, not more than twice longer than distance between them. Cupulae ia shaped as in males. Lateral setae c2, d2, e2, f2 short, piliform. Distances e1:e1 longer than h1:h1, e1:h1 three times longer than h1:h1, h1:h1 as long as the diameter of supranal concavity. Setae h2 shaped as macrosetae, h3 absent on the slide; ps1 hair-like, slightly longer than distance between them. Distance between dorsal setae: se:se 71, si:si 58, c1:c1 50, d1:d1 32, c1:d1 70, c2:c2 132, d2:d2 92, c2:d2 96, e1:e1 20, e2:e2 98, d1:e1 90, d2:e2 90, h1:h1 16, e1:h1 36, h2:h2 47, h3:h3 28, ps1:ps1 10, h1:ps1 33. Sclerotization of coxal fields similar to that in males, except for lateral longitudinal shields of epimerites IV devoid of medial shields. Opisthoventral sclerites absent. Epigynum shaped as a large horseshoe-like sclerite, width 48, length 66, L/W 1.4. Tips of the epigynum branches almost reach the level of setae g and extend beyond the ends of latigenital sclerites (Fig. 4). Genital acetabula set Szeptyckiana gen. n. from falconets 71

Fig. 6. Photos of male gnathosoma (A, B) and ventral hysterosoma (C, D). A, C – S. microsetosa sp. n., B, D – S. macrosetosa sp. n.

posterior to setae 3a and at the level of 3b on epigynum branches. Distance 3b:g similar to the dis- tance g:4a, 3a:3b shorterthan 3b:g. Legs IV reaching by ambulacra the level of setae ps2. Genual setae cG I, II dagger-like, approxi- matellyaslongasthewidthofcorrespondingpodomere(Fig.5E),longerthaninmales. Juvenilestagesunknown. J. DABERT, A. LABRZYCKA 72 E t y m o l o g y. The specific name (micro, L. – small + saeta, L. – hair) derives from the short, hair-likesetae vi. A c k n o w l e d g e m e n t s. We wish to thank Prof. Gerd ALBERTI, Zoological Institute and Museum, Ernst-Moritz-Arndt University, Greifswald, Germany, for making available samples of drybirdskinsforthepresentstudy.

REFERENCES

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